IMMUNOHISTOCHEMICAL STUDY OF ESTROGEN AND PROGESTERONE RECEPTORS IN DIFFERENT SIZE CLASSES OF OVARIAN FOLLICLES IN DROMEDARY CAMELS | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Assiut Veterinary Medical Journal | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Article 17, Volume 58, Issue 134, July 2012, Page 1-7 PDF (2.3 MB) | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Document Type: Research article | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
DOI: 10.21608/avmj.2012.173811 | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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Authors | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
A.M. SALEH1; E.A. ABDELHAFEZ1; D.R.I. DERAR2 | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
1Dept. of Anatomy and Histology, Fac. Vet. Med., Assiut University | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
2Dept. Theriogenology, Fac. Vet. Med., Assiut University | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Abstract | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
The present study aimed to investigate the normal distribution of estrogen (ERα) and progesterone (PR) receptors in the different classes of the ovarian follicles of the she-camel and their relation with the serum and follicular estrogen and progesterone hormonal level. Progesterone (P4) and 17β Estradiol (E2) serum levels, follicular estrogen receptors (Erα) and progesterone receptors (PR) expression and morphometry of different size classes of the ovarian follicles and corpus luteum during the estrous cycle in female dromedary camel were evaluated.The ERα and PR were detected using an indirect immunohistochemistry method (streptavidin-biotin immunoperoxidase method). Serum and follicular fluid were taken from 10 female dromedaries before and after slaughtering to evaluate the level of estrogen and progesterone in different size classes of the ovarian follicles (small >10mm, medium 10-15 and large <15 mm). Three cyclic non-lactating reproductively healthy she-camels were examined and bled daily to assess serum estrogen and progesterone levels during the follicular cycle and relate them to the size of the ovarian follicles and corpus luteum. The hormone levels were measured by radioimmunoassay. ERα was detected at low amounts in the follicular cells of the primordial, primary follicles and corpora lutea while, detected at moderate in the secondary follicles and high in the oocytes and in the tertiary follicles. On other hand, PR was detected in low reaction in secondary follicles and tertiary follicles. Moreover, it was estimated in a moderate reaction corpora lutea and corpora albicantia and stroma cells and in a strong reaction in the blood vessels. The number of cells expressing Erα and PR protein in ovarian cells was assessed by immunohistochemistry and computerized morphometric procedures. All data were analyzed with two way repeated measures ANOVA. Highest expression levels (p <0.01) of ER was found in the oolemma of the small and medium follicles but was absent in the granulosa cells surrounding these follicles. ERα was widely dispersed in the theca interna and theca externa of large tertiary follicles, it showed moderate staining intensity with homogeneous distribution in the cellular nucleus in theca interna and strong staining in the theca externa. The area and P4 serum concentration were higher during the luteal phase in R and C cows, as compared to F1. C and R animals displayed higher serum hormone levels than the F1 genotype, expression of mERα and glandular mPR and glandular sizes, which could pertain to the previously found better reproductive performance of C and R genotypes, probably as a result of their adaptation to a low-land tropical environmentExpression of PR was observed in stroma of all luteal structures but significantly stronger reaction was found in blood vessels of these structures. Estrogen concentration in both the follicular fluid (r=-0.06) and serum (r=-0.15) correlated negatively -but (not significantly)- with the size of the follicle while a positive non significant correlation was found between serum progesterone and the size of the corpus luteum. Serum and follicular fluid estrogen was higher in follicles exceeding 15 mm more than the lesser follicular categories. Slight difference in the concenttration of estrogen was found between follicles less than 10 mm in diameter and those between 10 – 15 mm. The expression of ER and PR and the secretion of their specific hormones in the ovary of she-camel were not always correlated with the presence of the hormones. | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Keywords | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
Key words: Immunohistochemistry; ER; PR; Camel ovary | |||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||||
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IMMUNOHISTOCHEMICAL STUDY
A.M. Saleh *; E.A. Abdelhafez* and D.R.I. Derar ** * Dept. of Anatomy and Histology, Fac. Vet. Med., Assiut University ** Dept. Theriogenology, Fac. Vet. Med., Assiut University
__________________________________________________________________________________________ ABSTRACT __________________________________________________________________________________________
__________________________________________________________________________________________ Key words: Immunohistochemistry, ER, PR, Camel ovary. __________________________________________________________________________________
INTRODUCTION
Steroid hormones are important regulators of reproductive processes in female mammals. Estrogens and progesterone receptors mediate respectively the action of estrogens and progesterone by regulating transcription target genes. Estrogens possess an intrafollicular action by stimulating in synergic manner with FSH the aromatase activity (Adashi et al., 1982 and Fitzpatrick
Receptors for estrogen (ER) are expressed as 2 structurally related subtypes in mammals, ERα and ERβ, which are encoded by 2 distinct genes. The existence of these 2 subtypes may partly explain the selective action of estrogen in different target tissues and in the same tissue in different physiological statuses (Conneely,
MATERIALS and METHODS
Collection of samples Ovaries of 35 adult female one-humped camels (Camelus dromedarius) slaughtered at a Cairo abattoir, were collected immediately after slaughter. The period from November to April was taken as the peak breeding season, while May-October was considered as the low breeding season.
Pre-slaughter information regarding the nutritional or reproductive status of these camels was not available. After cleaning each ovary off the extraneous tissue, diameter of Graafian follicles was measured using Vernier Calipers. On the basis of their size, follicles were classified into three groups viz. small (5-9 mm) and large (10-20 mm). Fluid from each follicle was aspirated aseptically and stored at –20°C. Animals having ovaries with any pathological lesions, or those with cystic follicles (>20 mm in diameter; Tibary and Anouassi, 1996) were not included in the study. Before slaughter, about 15 ml peripheral blood was collected from each animal, serum was separated and stored at –20°C for hormonal analysis.
Histological and immunohistochemsitry:
Blood serum and follicular fluid samples were analyzed for progesterone, estrogen, through EIA technique, using a Microstrip Elisa Reader (Stat-Fax-303, Awareness Technology, Inc.). Progesterone and estradiol concentrations were determined by using kits from Bremancos Diagnostic INC–GmbH, Germany (Cat. # BC-1113 & BC-1111, respectively). The lowest detectable level of progesterone during this test was 0.05 ng/ml, while the cross reactivity with other steroid hormones was <0.74%. For estradiol, the lowest detectable level was 5.9 pg/ml and cross reactivity with other steroids was <2.10%.
Statistical analysis: Statistical analysis of the collected data was carried out according to procedures of completely random design, SAS (1995).
RESULTS
Morphologically, the ovary of the she-camel was flattened, lobulated measuring 3.17, 2.21 and 0.8 cm length, width and thickness respectively. Both right and left ovaries exhibited follicles in various stages of development, including primordial, primary, secondary and tertiary follicles, corpora albicantia, and late CL, as well as follicles with different degrees of atresia. The measurements of the different follicles and corpora lut
Immunohistochemically: In all examined she- camels,
In mature ovarian follicles (Fig. 4); ERα was strongly expressed in the cellular nuclei and cytoplasm of the granulose, and moderate in that of theca interna, and theca externa layers.
On other hand,
Serum and follicular fluid estrogen (Table. 2& 3
Table 1: The morphmetric measurements of the oocytes in µm in the different ovarian follicles measured from histological sections of the she-camel ovaries stained with H&E
Table 2: Estrogen (E2) concentration in follicular fluid and serum of different size classes of ovarian follicles of she camels.
Table 3: Progesterone concentration in she camels relative to the size of the CL
Hist. (1): Estrogen (E2) concentration in follicular fluid and serum of different size classes of ovarian follicles of she camels.
Hist. (2): progesterone concentration in she camels relative to the size of the CL
1 2
Fig. 1- 4: Micrographs of different she-camel ovarian cell types at various estrous stages showing immunostaining of ERα. By Streptavidin-Biotin method, Mayer's hematoxylin counterstain.
1- Outer region of the ovarian cortex during estrus with high immunoreactivity for ERα (SERα) in the surface epithelium, low immunoreactivty in the superficial stroma, and strong expression in the tunica albuginea, cytoplasm and nuclus of oocytes of the primordial follicles.X200.
2- Primary follicle of she-camel ovary showed ERα strong immunoreactions in the cytoplasm of oocyte and moderate reaction in the follicular cells. X100.
3- Low ERα immunostaining in the wall of a secondary follicle (s), stroma cells and atric follicles during proestrus. X100 .
4- Wall of the mature ovarian follicle presented strong immunoreactivity in the cytoplasm and nuclei granulosa, whereas theca interna and externa show moderate reaction but few nuclei show strong reaction. X200.
Fig. 5-6: Micrographs of different she-camel ovarian cell types showing immunostaining of PR. By Streptavidin-Biotin method, Mayer's hematoxylin counterstain.
5- PR immunoreactivity was moderated in corpus luteum, cells of the stroma and blood vessels. X40.
6- In the mature ovarian follicle, the PR immunoreactivity was low in the theca interna and externa but high in granulose layer X40
DISCUSSION
The obtained immunohistochemical observations indicate that the expression of ER and PR and the secretion of their specific hormones in the ovary of Arabian she-camel were well correlated with the reproductive cycle. But during ovarian activity, the expression of ER and PR is not always correlated with the presence of the hormones in the follicles and serum. The immunohistochemical expression of the ER was detected in the nuculei and cytoplasm of the ovarian cells whereas in the primate ovaries, the ER of the granulosa cells was nuclear only (Billiar et al., 1992; Suzuki et al., 1994 and Saunders et al., 2000). In rat, hamster and pig, the ER is nuclear and cytosolic, and the cytosolic fraction is more important (Kawashima and Greenwald 1993). According to Guiochon-Mantel and Milgrom (1993), ER is essentially localized in the nucleus in the absence of estrogens. In addition, at the hypothalamic and hypophysal level, the estradiol injection induces the increase of the cytosolic fraction of the ER (Kawashima et al., 1987). In she-camel ovary, the variation during the reproductive cycle would translate a functional action. Numerous studies show on the contrary that the theca cells are the major sites of ER expression in different speciesincluding rats, mice and primate (Chandrasekher et al., 1994; Kuiper, et al., 1996; Sanchez-Criado et al., 2005) or total absence of the ER in the ovary (Saunders et al., 1997).
As reported in the bovine ovaries (Van den Broeck et al., 2002), in the present study, the PR localized in the different ovarian structure of the she-camel additionally shows that the various ovarian cell types exhibit different patterns of PR immunoreactivity during the ovarian activities. In the follicle cells of primordial, primary and secondary follicles the scores for PR were high and increased from primordial to secondary follicles. These data are in accordance with findings in primates (Hild-Petito et al., 1988) and dogs (Vermeirsch et al., 2001), and they indicate that progesterone may regulate follicular growth during the early stages of follicular development.
In she-camel follicular structures examined, the follicle/granulosa cells showed the high PR immunostaining during oestrus, when ovulation occurs. These results are concomitant with earlier observations in the dog (Vermeirsch et al., 2001) and with a study on PR mRNA in the bovine ovary (Cassar et al., 2002). The crucial role of PR in the ovulatory process has been demonstrated in PR-deficient mice, since such mice develop large follicles but fail to ovulate (Lydon et al., 1995). All these findings emphasize the important role of progesterone and its receptor in the ovulation process. The expression of PR in granulosa cells in tertiary follicles is induced by the LH surge (Hild-Petito et al., 1988). The induction of PR mRNA has also been observed in monkey granulosa cells during periovulatory stages (Chandrasekher et al., 1994) and in porcine granulosa cells cultured in vitro after LH stimulation (Iwai et al., 1991). The progesterone receptors are reported to mediate the protective effects of progesterone against apoptosis in the granulosa cells of preovulatory follicles (Quirk et al., 2004).
The presence of PR in corpora lutea reflects the role of progesterone in corpus luteum activity (Revelli et al., 1996; Rueda et al., 2000). Progesterone regulates the proliferation and development of luteinized granulosa and theca cells in an autocrine and paracrine way (Sasano and Suzuki, 1997). The presence of PR in all lutein cells of the corpora lutea suggests the influence of progesterone in the luteinization process (Revelli et al., 1996; Duffy et al., 1997). However, in the present study, the PR immunostaining in the corpus luteum was lower than in most other ovarian structures, which can be due to a negative effect of the locally produced high levels of progesterone to the PR production. In contrast to all other ovarian cells, the lutein cells of the corpora lutea showed PR immunostaining not only in the nuclei, but in the cytoplasm as well.
A low but manifest PR immunoreactivity was observed in cells of the tunica albuginea and the surface epithelium. This corresponds with a study on ovine ovaries in which it has been suggested that cells of the ovarian surface epithelium are enzymatically involved in the ovulation process by the influence of progesterone and its receptors (Murdoch, 1998). Further investigations in cattle are necessary to verify the role of PR and progesterone in the ovarian surface epithelium.
The present study indicated that estrogen concentration in both the follicular fluid (r = 0.06) and serum (r = 0.15) correlated negatively (non-significantly) with the size of the follicle while a positive non significant correlation was found between serum progesterone and the size of the corpus luteum. However, positive non significant correlation was detected between serum progesterone and the size of the corpus luteum in the studied animals throughout the days of the cycle (r = 0.25). On other hand, present immunohistochemical observations indicate that the expression of ER and PR and the secretion of their specific hormones in the ovary of camel was not always correlated with the presence of the hormones.
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Chandrasekher, YA.; Melner, MH.; Nagalla, SR. and Stouffer, RL. (1994): Progesterone receptor, but not estradiol receptor, messenger ribonucleic acid is expressed in luteinizing granulosa cells and the corpus luteum in rhesus monkeys. Endocrinol., 135: 307-314. Conneely, OM. (2001): Perspective: female steroid hormone action. Endocrinol., 142: 2194–2199, Duffy, DM.; Wells, TR.; Haluska, GJ. and Stouffer, RL. (1997): The ratio of progesterone receptor isoforms changes in the monkey corpus luteum during the luteal phase of the menstrual cycle. Biol. Reprod.; 57: 693-699. Fitzpatrick, SL. and Richards, JS. (1992): Regulation of cytochrome P450 aromatase messenger ribonucleic acid and activity by steroids and gonadotropins in rat granulosa cells. Endocrinol., 129(3): 1452-1462. Graham, JD. and Clarke, CL. (1997): Physiological action of progesterone in target tissues. Endocrinol. Rev., 18(4): 502-519. Guiochon-Mantel, A. and Milgrom, E. (1993): Cytoplasmic-nuclear trafficking of steroid hormone receptors. Trends Endocrinol. Metabol., 4: 322-328. Hild-Petito, S.; Stouffer, RL. and Bremmer, RM. (1988): Immunocytochemical localization of estradiol and progesterone receptors in the monkey ovary throughout the menstrual cycle. Endocrinol., 123: 2896-2905. Iwai, M.; Yasuda, K.; Fukuoka, M.; Iwai, T.; Takakura, K.; Taii, S.; Nakanishi, S. and Mori, T. (1991): Luteinizing hormone induces progesterone receptor gene expression in cultured porcine granulosa cells. Endocrinol., 129: 1621-1627. Kawashima, M. and Greenwald, G. (1993): Comparison of follicular estrogen receptors in rat, hamster and pig. Biol. Reprod., 48: 172-179. Kawashima, M.; Kamiyoshi, M. and Tanaka, K. (1987): Presence of estrogen receptors in the hen hypothalamus and pituitary. Endocrinol., 120: 582-588. Kogo, H.; Fujimoto, T. and Mori, T. (1999): Evidence for gonadotropin releasing hormone receptor mRNA expression by estrogen in rat granulosa cells. Cell Tissue Res., 297: 459-465. Kuiper, GGJM.; Enmark, E.; Pelto-Huikko, M.; Nilsson, S. and Gustafsson, JA. (1996): Cloning of a novel estrogen receptor expressed in rat prostate and ovary. Proc .Nalt. Acad. Sci., 93: 5925-5930. Lydon, JP.; DeMayo, FJ.; Funk, CR.; Mani, SK.; Hughes, AR.; Jr Montgomery, CA.; Shyamala, G.; Conneely, OM. and O'Malley, BW. (1995): Mice lacking progesterone receptor exhibit pleiotropic reproductive abnormalities. Genes & Development., 9: 2266-2278. Murdoch, WJ. (1998): Perturbation of sheep ovarian surface epithelial cells by ovulation: evidence for roles of progesterone and poly (ADP-ribose) polymerase in the restoration of DNA integrity. J. Endocrino., 157: 503-508. Nomina Histologica (1994): International Committee on Veterinary Histological Nomenclature, revised 2nd ed., p. 38. Cornell, NY, Palter, SF.; Tavares, AB.; Hourvitz, A.; Veldhuis, JD. and Adashi, EY. (2001): Are estrogens of import to primate/human ovarian folliculogenesis? 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دراسة کيميائية هستومناعيه علىمستقبلاتهرمونالاستروجينوالبروجستيرونفيجريبات المبيض المختلفة في الجمال العربية
عبد المهيمن مصطفى صالح ،ايناس احمد عبدالحافظ ،ضرار رفعت ضرار
تهدف هذه الدراسة إلى دراسة توزيع مستقبلات هرمون الاستروجين والبروجستيرون في مختلفة جريبات المبيض من الناقة وعلاقتها مع مستوى هرمون الاستروجين والبروجسترون فى السائل الجريبى والدم. تم دراسة مستبقبلات الاستروجين والبروحستيرون بواسطة طريقة تفاعل کيمياء النسيج المناعى الغير مباشرة. وتم قياس مستويات الهرمونات في الدم بواسطة المقايسة المناعية الاشعاعية. وجدت مستقبلات الاستروجين في کميات قليلة في الخلايا الجريبية لجريبات المبيضية الأساسية والاولية وکذلک فى الجسم الاصفر في حين وجدت متوسطة في الخلايا الجريبية للجريبات الثانوية وعالية فى البويضات والخلايا الجريبية للجريبات المبيضية الناضجة. في المقابل وجدت مستقبلات البروجستيرون فى کميات منخفضة في الجريبات المبيضية الثانوية والناضجة، ومعتدلة التفاعل فى الأجسام الصفراء وعالية التفاعل في الأوعية الدموية. وجد ان العلاقة بين ترکيزهرمون الاستروجين في کل من السائل الجريبي ومصل الدم مع حجم الجريبات کانت عکسية، ولکن ليست بشکل کبير، في حين وجد ارتباط إيجابي کبير بين ترکيز البروجسترون في مصل الدم وحجم الجسم الأصفر. وکما وجد ان ترکيز الاستروجين فى مصل الدم والسائل الجريبي کان عاليا في الجريبات الاکبر من 15 مم أکثر من الجريبات الأقل حجما. لکن الاختلاف کان طفيفا بالجريبات المبيضية الأقل من القطر 10مم وتلک ما بين 10 - 15 مم. کما وجد ان توزيع مستقبلات هرمون الاستروجين والبروجستيرون المبيضى غير مرتبط بمستوى الهرمونين فى الدم والسائل الجريبى.
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REFERENCES Adashi, EY. and Hsueh, AJW. (1982): Estrogens augment the stimulation of ovarian aromatase activity by follicle stimulating hormone in cultured Rat granulosa cells. J. Biol. Chemistr., 257:6077-6083.
Amrozi, Kamimura, S.; Ando, T. and Hamana, K. (2004): Distribution of estrogen receptor α in the dominant follicles and corpus luteum at the three stages of estrous cycle in Japanese black cows. J. Vet. Med. Sci., 66: 1183–1188.
Berisha, B.; Pfaffl, M. and Schams, D. (2002): Expression of estrogen and progesterone receptors in the bovine ovary during estrous cycle and pregnancy. Endocrinol., 17: 207–214.
Billiar, RB.; Loukides, JA. and Miller, MM. (1992): Evidence of the presence of estrogen receptor in the ovary of baboon (Papio anubis). J. Clin Endocrinol. Metabol., 75 (4): 1159-1165.
Cassar, CA.; Dow, MPD.; Pursley, JR. and Smith, GW. (2002): Effect of the preovulatory LH surge on bovine follicular progesterone receptor mRNA expression. Domestic Anim. Endocrinol., 22: 179-187.
Chandrasekher, YA.; Melner, MH.; Nagalla, SR. and Stouffer, RL. (1994): Progesterone receptor, but not estradiol receptor, messenger ribonucleic acid is expressed in luteinizing granulosa cells and the corpus luteum in rhesus monkeys. Endocrinol., 135: 307-314.
Conneely, OM. (2001): Perspective: female steroid hormone action. Endocrinol., 142: 2194–2199,
Duffy, DM.; Wells, TR.; Haluska, GJ. and Stouffer, RL. (1997): The ratio of progesterone receptor isoforms changes in the monkey corpus luteum during the luteal phase of the menstrual cycle. Biol. Reprod.; 57: 693-699.
Fitzpatrick, SL. and Richards, JS. (1992): Regulation of cytochrome P450 aromatase messenger ribonucleic acid and activity by steroids and gonadotropins in rat granulosa cells. Endocrinol., 129(3): 1452-1462.
Graham, JD. and Clarke, CL. (1997): Physiological action of progesterone in target tissues. Endocrinol. Rev., 18(4): 502-519.
Guiochon-Mantel, A. and Milgrom, E. (1993): Cytoplasmic-nuclear trafficking of steroid hormone receptors. Trends Endocrinol. Metabol., 4: 322-328.
Hild-Petito, S.; Stouffer, RL. and Bremmer, RM. (1988): Immunocytochemical localization of estradiol and progesterone receptors in the monkey ovary throughout the menstrual cycle. Endocrinol., 123: 2896-2905.
Iwai, M.; Yasuda, K.; Fukuoka, M.; Iwai, T.; Takakura, K.; Taii, S.; Nakanishi, S. and Mori, T. (1991): Luteinizing hormone induces progesterone receptor gene expression in cultured porcine granulosa cells. Endocrinol., 129: 1621-1627.
Kawashima, M. and Greenwald, G. (1993): Comparison of follicular estrogen receptors in rat, hamster and pig. Biol. Reprod., 48: 172-179.
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